Historia naturalis bulgarica, 9, 1998: 53-69

Fulica atra pontica subsp. n, from the Middle

Holocene on the South Black Sea Coast, Bulgaria

Zlatozar BOEV, Elena KARAIVANOVA

Introduction

The coot is represented in the recent avifauna of Bulgaria by the nominate

subspecies Fulica atra atra Linnaeus, 1758, spread in the Palearctic up to the

Hindustan Peninsula.

Eleven recent species have been included in the genus Fulica Linnaeus, 1758.

America is considered the center of speciation, where 9 species are distributed, 6 of

them endemic for the southern and western regions of South America (Taylor, 1996).

Description of the site

The site (UTM code NG 59) is a submerged prehistoric settlement in the

Sozopol Bay (the Bulgarian Black Sea Coast), NE of the town of Sozopol, nowsubmerged at 12 m depth.

The site is dated back to the end of the Eneolithic period (end of 7th - early

6th millenium B.P.) - Early Bronze Age (4800-4400 B.P.) (Spassov & Iliev, 1995). It

was studied in the underwater excavations, organized by Dr Mihail Lazarov

(1987-1990) and the archaeologists Dr VesseHn Draganov and Dr Hristina

Angelova. We received the avian material from Dr Lazar Ninov, Dr Georgi

Ribarov, Dr Nikolay Spassov and Nikolay Iliev. N. Spassov has studied all the

mammalian bone remains of the site.

Associated fauna. Gavia arctica, Podiceps cristatus, Phalacrocorax carbo,

Ardea cinerea, Anas querquedula, Anas platyrhynchos, Aythya nyroca, Fulica

atra (the last species was the most numerous game and it constituted over 52 %of the gamefowl after BOEV, 1995a); Lepus capensis, Bos primigenius, Cervus ela-

phus, C. dama, Capreolus capreolus, Sus scrofa, Felis silvestris, Vulpes vulpes,

Meles meles, Equus gmelini, Panthera leo, Tursiops truncatus, Delphinus del-

phis, Phocaena phocaena, Monachus monachus, Thunnus thunnus, Scomber

53

scombrus, Cyprinus carpio, Testudo graeca and Emys orbicularis. Among the

domestic animals are: Bos taurus, Ovis aries, Capra hircus, Sus scrofa domestica,

Canus familiaris and Equus caballus (RiBAROV, 1991; Spassov & Iliev, 1995; G.

Ribarov - impubl. data).

Climatic data: The climate of the region is temperate. The average annual

temperature is 13,3° and the average annual amplitude of temperature is 20,6°

The precipitation maximum is in autumn, while the precipitation minimum is

in summer. The winter in the region is the warmest one for the country and the

January average temperature is + 3,2° (Mishev et al, 1989).

Ecobiogeographical characterization of Fulica atra

Fulica atra is a partly resident, partly migratory species. It nests from the

boreal to the subtropical zone in larger water bodies, avoiding freezing waters. It

winters in open parts of the sea shore (Harrison, 1982). The coot is the most

aquatic of all West-Palearctic rails. It prefers shallow water for diving and suffi-

cient space giving the opportunity for running on the water surface in the taking

off. It tolerates the wind and the waves in the sea. It is considered a lowland

species, but nevertheless it inhabits suitable habitats up to 1100 m a.s.l. in the

Alps (Cramp & Simmons, 1980). An ancient species of Miocene (i.e. of Pliocene, fol-

lowing more recent views) origin. It has spread northward and northeastward

from the southern coastal regions of Eurasia after the Pleistocene

(VOINSTVENSKIY, 1960).

A common and numerous resident (in South Bulgaria), breeding migratory

species. Inhabits both salt and fresh wetlands in the lowlands and plains through-

out the country. The Balkan population prefers water basins with dense aquatic

vegetation. Before the drastic destruction of the suitable habitats along the

Danube River banks and the Black Sea shore it was still numerous and widely

used as gamefowl (MiCHEV, 1990).

Fossil and subfossil record of Fulica atra

Several dozens of quaternary localities of the species are known from the

world literature. They were summarized for the first time by Brodkorb (1967),

who reported on the Pleistocene-Holocene distribution of F. atra in Ireland,

England, Wales, Italy, Switzerland, and Azerbaijan.

Many data for the Quaternary finds of the species may be drawn from the

more recent literature:

Pleistocene: the middle and lower streches of the rivers Dniepr, Don and Ural

(VOINSTVENSKIY, I960); from the interstadial Wurm 2-3 to the Atlantic period -

54

25 340 to 10 830 .. in the Shandaja Cave in Croatia (Malez-Bacic, 1979);

Epipaleolithic (Natufian) of Mallaha in Palestine (Pichon, 1987); Middle

Pleistocene in Prezletice in Czechia (Janossy, 1983a); Lower Pleistocene in the

Velika Pecina Cave in Croatia (Malez-Bacic, 1975); Middle-Late Villafranchian in

the Sandalija 1 Cave and of Late Pleistocene (Wurm 2) in the Sandalija 2 Cave in

Croatia (Malez-Bacic, 1979); La Page in France, Grimaldi in Italy and Istallosko

in Hungary (Malez-Bacic, 1979); Late Pleistocene and Eneolithic in the Velika

Pecina Cave in Croatia (Malez, 1975); Pleistocene of Crete (Weesie, 1988); Early

Pleistocene in Norfolk in England (Harrison, 1985); Wurm 3 (28 000 B.P.) to

Postglacial (ca. 4000 B.P.) in a number of sites in S France and Catalonia in Spain

(Vilette, 1983); Late Paleolithic (14 570 - 11 380 B.P.) in Cueva de Nerja in Spain

(Hernandez, 1995); Late Pleniglacial ca. 33 000 B.P. in the Oblazowej Cave in

Poland (TOMEK & BOCHENSKI, 1995); Middle and Late Pleistocene of Corsica and

Crete (Alcover et al, 1992); fmal of Wurm 3 (20 000 B.P. - middle of Wurm 4 (12

500 B.P.) in Arene Candide in S Italy (Cassoli, 1980); Late Pleistocene in Grotta

di Cala Genovesi, Levanzo (Cassoli & Tagliacozzo, 1982) and Grotta della

Madonna, Calabria in Italy (Cassoli, 1992) and Paleolithic in the Alimovskiy

Naves Cave in Crimea (Baryshnikov & Potapova, 1992).

Holocene: Holocene of middle and lower streches of the rivers Dniepr, Don and

Ural (Voinstvenskiy, 1960); the Russian town of Voin' in the Middle Ages in

Poltava Region (Sergeev, 1965); the settlement of Russeshti from the 4th milleni-

um B.C. (Ganya, 1972); Subatlantic period in Baile 1 Mai in Romania (Kessler,

1985); Late Pleistocene in the lower Nil Valley in Egypt (Gautier, 1988); Early

Paleolithic of Grotta dei Fanciulli in Balzi Rossi in Italy (Campana, 1946); Late

Pleistocene in Torre in Pietra in Italy (Cassoli, 1978); 1350-1520 A.D. in London

(Bramwell, 1975);

Quaternary finds of Fulica atra in Bulgaria

Besides the locality of Sozopol, 6 other Quaternary sites (Fig. 1) of: Fulica atra ore

known in Bulgaria (Boev, 1996 a, b; 1997). AU finds originate from the Holocene

deposits and are kept at the National Museum of Natural History in Sofia (NMNHS):- Urdoviza near the village of Kiten, Burgas Region. Early Bronze Age (3000-

2000 B.C.): humerus dex., NMNHS 1099; humerus dex. - NMNHS 4416 (Boev &Ribarov, 1990);

- Yayla near Albena Resort, Dobrich Region. ? Middle Holocene: ulna sin.

prox., NMNHS 631;

- Topchii Village, Razgrad Region. Early Holocene: carpometacarpus sin.,

NMNHS 1083; tibiotarsus dex. dist., NMNHS 1087; tbt dex. dist., NMNHS 3123;

cmc dex. dist., NMNHS 3125.

- Durankulak, Dobrich Region. Late Holocene (1-4 century A.D.): humerus sin.,

55

Ji-—"=-==-=—=-"-=

Table 1

Measurements of the skeleton of recent (R) and fossil (F) Fulica atra from SEBulgaria

N of the feature R/

F

N x SD min max

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

21

22

23

57

R

Table 1 (continuation)

N of the feature R/ F N SD

24

25

26

27

28

29

30

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

46

F

Table 1 (continuation)

N of the feature

ter of the radius proximal epiphysis (31); smaller diameter of the radius proximal

epiphysis (32); width of the distal radius epiphysis (33); height of the distal radius

epiphysis (34); total length of the femur (35); width in the middle of the femur dia-

physis (36); trochanter major diameter (37); minimum width of the collum femoris

(38); diameter of the caput femori (39); width of the femur proximal epiphysis

(40); diameter of the condylus fibularis (41); maximum width of the distal femur

epiphysis (42); diameter of the femur condylus medialis (43); diameter of the

femur condylus lateralis (44); total length of the tibiotarsus (45); width in the mid-

dle of the tibiotarsus diaphysis (46); minimum width of the tibiotarsus diaphysis

(47); width of the distal tibiotarsus epiphysis (48); minimum width of the proxi-

mal tibiotarsus epiphysis (49); maximum width of the proximal tibiotarsus epiph-

ysis (50); length of the proximal tibiotarsus epiphysis (51); diameter of the tibio-

tarsus condylus medialis (52); diameter of the tibiotarsus condylus lateralis (53);

minimum width of the trochlea tibiotarsi (54); maximum width of the tibiotarsus

distal epiphysis (55); total length of the tarsometatarsus (56); width in the middle

of the tarsometatarsus diaphysis (57); minimum width of the tarsometatarsus

diaphysis (58); width of the tarsometatarsus distal epiphysis (59); width of the

trochlea 3 of the tarsometatarsus distal epiphysis (60); width of the tar-

sometatarsus proximal epiphysis (61); height of the tarsometatarsus proximal

epiphysis (62); height of the tarsometatarsus fovea medialis (63); diameter of the

tarsometatarsus trochlea 3 (64); diameter of the tarsometatarsus trochlea IV (65).

The Eneolithic population of Fulica atra

from the Sozopol Bay

Fulica atra pontica subsp. n.

Holotype: Complete tibiotarsus dex., collection

of the NMNHS, No 6568, collected in 1987 by Dr

Georgi Ribarov (Fig. 2).

Paratypes: Topotypes, NoNo 1147-1175; 6566-

6567; 6569-6595; 6597-6628; 6630; 6632-6634; 6636-

6637; 6641; 6642; 6644-6655; 6658-6701; 6704; 6706-

6708; 6713, collected by Dr G. Ribarov in 1987-1990

and Dr Nikolay Spassov. See Table 1 (column „n")

and the descriptions of the measurements in

„Material and Methods" for the distribution of the

paratypes in anatomical elements.

Locality: Submerged settlement at 12 m depth

in the Sozopol Bay (Bulgarian S Black Sea Coast).

Fig. 2. Fulica atra pontica ssp. n. (holotype, NMNHS 6568): medial view (left) and cranial

view (right) (Photographs: Boris Andreev)

60

f IL_L

Fig. 3. The manner of measurings of the skeletal elements of Fulica atra: a - sternum, b -

coracoid, - scapiila, d - humerus, - ulna, f - radius (Drawings: Vera Hristova)

61

64 ^«^r^>

Fig. 3. The manner of measurings of the skeletal elements of Fulica atra: g - femur, h

tibiotarsus, i - tarsometatarsus (Drawings: Vera Hristova)

62

Horizon: Middle Holocene stratified sandy sediments, containing fossils of

hunted animals and tools of the ancient inhabitants of the settlement.

Chronology: Middle Holocene (end of the Atlantic period and the Subboreal

period): final of Eneolithic (end of 5th - early 4th millenium B.C.) to Early Bronze

Age (2800-2400 B.C.).

Etymology: The name pontica is given for the ancient Greek name of the

Black Sea - Pontes, whitch SW coast the subspecies remains originated from.

Measurements: See Table 1, Fig. 3.

Comparison: The general shape of all skeletal elements unambiguously

relates the finds to the family Redlidae and more exactly to its larger recent rep-

resentatives. Metrically and morphologically Fulica atra pontica ssp, n. is very

close to the recent Palearctic nominant subspecies F. a. atra. Porzana, Rallus,

Crex, Gallinula and Porphyrula are much smaller, while Porphyria has larger

dimensions and shows considerable differences in the bone structures.

Over one fourth of the osteometrical features (27,7 %), i.e. 18 of all the 65 stud-

ied features show significant statistical differences between the sizes of fossil and

recent coots. Four features show a very high and significant degree of guarantee

of the differences. In two of them it is 0,99, and in two others - 0,98 (Table 2).

Eight features are statistically different with guarantee of 0,95, and two others -

in 0,9.

Diagnosis: An extinct Middle Holocene subspecies of coot, which hsis statis-

tically significant differences (larger dimensions) in comparison with F. a. atra in

Table 2

Means of the osteometrical features of recent (r) and fossil (f) Fulica atra fromSE Bulgaria

N of the feature

Table 2 (continuation)

N of the feature

relation to the minimum width of the tibiotarsus diaphysis (t = -4.01904), the

minimum diameter of the distal epiphysys of the tibiotarsus (t= -3.10276), the

maximum width of the articular end of the scapula (t = -3.05487), and the diam-

eter of the condylus fibularis of the femur (t = -2.45676).

Comparative Material Examined: The find was compared with analogous

skeletal elements of the following species of the collection of NMNHS: Gallinula

chloropus- 4/1983, 3/1983, 10/1990, 11/1990; Fulica atra- 4/1986; 5/1986; 6/1986;

7/1986; 8/1986; 9/1986; 10/1986; 11/1986; 12/1986; 14/1987; 15/1989, 16/1989,

17/1990; 21/1996; Porphyrio porphyria - 1/1989, Rallus aquaticus - 1/1990,

2/1990, 3/1993, 4/1994; Crexcrex- 1/1986, Porzana porzana - 1/1989.

Discussion: Olson (1977) reported about 2 fossil subspecies of Fulica: F.

chathamensis chathamensis Forbes, 1892 and F. ch. prisca Hamilton, 1893 from

the Quaternary on the Chatham Island and the South Island of New Zealand. F.

hestera is considered as a synonym of F. americana. No fossil subspecies have

been described for F. atra.

The bone system is the most conservative structure (except the teeth) of the

living body. In vertebrates it reflects more completely the influence of the envi-

ronmental factors. Thus, the presence of 17 features of statistically significant

differences in the size dimensions in both samples (recent and fossil), allows to

conclude that the established differences show a taxonomical difference.

Therefore, the diferentiation of the fossil population of the submerged settlement

in the Sozopol Bay is completely reasonable.

The description of new taxa (subspecies and species) from the Quaternary

deposits in paleornithology is made on the base of small, scarcely perceptible, but

stable differences in the skeletal morphology (Janossy, 1987). Mourer-Chauvire

(1975a), for example, described Aquila chrysaetos binifacti on the basis of 18 fea-

tures (from a total of 89) of statistically significant differences from the Middle

Pleistocene in France. The reasons for the description of Buteo rufinus jansoni

(Mourer-Chauvire, 1975b) are similar.

As seen from Table 2 the dimensions of fossil specimens are statistically larg-

er for 18 features: P = 0,001 (feature No 47); P = 0,01 (No 44, 54); P = 0,02 (No 41,

48); P = 0,05 (No 4, 9, 11, 18, 30, 42, 43, 56, 57), and P = 0,1 (No 55, 58, 59, 64).

The metrical differences which speak to the advantage of fossil population

concern the elements of the fore girdle (scapula, coracoid and humerus), and the

main three long bones of the hind girdle (femur, tarsometatarsus and tibiotar-

sus), of the other hand. They may reflect some (possibly very small), differences

in the relative loading of the fore and hind limbs in the locomotion activity of the

birds. The more strongly developed articular surface of the scapula may be relat-

ed to the necessity of shorthening of the take-off way on the water surface. This

morphological adaptation correlates with the stronger development of the femur

and the tibiotarsus bones. It is quite possible to explane these morphological dif-

ferences by the ecological conditions, as for example by the presence of the more

65

densely vegetated water bodies, the longer stay and locomotion in the water, as

compared to these on the hard ground, etc.

The Pontian (the Black Sea) Coot possibly survived up to the Middle Holocene,

since when the proportional modification of the skeleton had been taking place

simultaneously with the emergence of the modern climate, gradually transform-

ing it into the recent subspecies. It is possible that the Sozopol fossil population

belonged to the winter migrants from the northeast, where the last influences of

the glaciation disappeared much later than on the southern parts of the Balkan

Peninsula. Part of the recent Baltic population at present winters in the Balkans.

On the other hand, the major winter concentrations in the Black Sea are sup-

posed to have originated from the birds of the western regions of the former

USSR (Cramp & Simmons, 1980). The hypothesis of a relict migration of a north-

ern population of coots in the Middle Holocene should not be rejected as a whole.

Unfortunately, we have not any comparative osteological material from the

northern parts of the species' range to evaluate such suppositions.

Acknowledgements

The authors express their thanks to Dr Georgi Ribarov (Historical Museum of

Burgas) and Dr Nikolay Spassov (NMNHS), who had lended us this material for

examination. We are also grateful to Dr Cecile Mourer-Chauvire (Universite

Claude Bernard - Lyon 1) for her helpful comments on the achieved results.

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Received on 8. 12. 1997

Author's address:

Dr Zlatozar Boev

National Museum of Natural ffistory

1, Tzar Osvoboditel Blvd

1000 Sofia, Bulgaria

Elena Karaivanova

Institute of Zoology

1, Tzar Osvoboditel Blvd

1000 Sofia, Bulgaria

68

Fulica atra pontica subsp. n. om,, I^PAMBAHOBA()

( u ):( 5 - 4. ...) go-(2800-2400 . ...) 159 .

(27,7 %), .. 18

65,u .

(0,999 go 0,98) - 1 0,999, 2 0,99 u

2 0,98. 9

0,95 u 4 - 0,90.

Fulica atra pontica ssp. .:, F. . atra (-)(t = -4.01904),

(t = -3.10276), condylus fibularis

(t= -2.45676) u (t = -2.55808).

F. . pontica ssp. ., go ,, u .u, ..

3UMHU ,-, .,,1" ( ) . moBa, kpamkompaUHOcm( 6000 .), ,Fulica atra..

69